Links for Palaeobotanists

Home / Introductions to both Fossil and Recent Plant Taxa / Angiosperms

Cyanobacteria and Stromatolites
Seed Plants in General
! Herbaria@
! Parasitic Plants@
! Taxonomy and Plant Classification Databases@
Plant Photographs@
Image Collections@
Picture Search@


A. Antonelli et al. (2015): An engine for global plant diversity: highest evolutionary turnover and emigration in the American tropics. In PDF, Front. Genet., 6.

! APG IV (2016): An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG IV. Open access, Botanical Journal of the Linnean Society, 181: 1–20.

! APG III (2009), compiled by B. Bremer et al.: An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG III. In PDF, Botanical Journal of the Linnean Society, 161: 105–121. See also here.

Hank Art et al., Williams College, Biology Dept., Williamstown MA: Field botany. Go to: Evolutionary Botany. Powerpoint download and links to aricles. See especially:
Early Land Plants.
Fossil Angiosperms.
Introduction to the Angiosperms.
Powerpoint presentations.

! Lorna Ash & Heather Kroening, Department of Biological Sciences, University of Alberta: Instructional Multimedia, Multimedia Topics, Botany. Snapshot taken by the Internet Archive´s Wayback Machine. Go to: Life Cycle of an Angiosperm. See also here. Online and downloadable flash 4 movies. Excellent!

B.A. Atkinson (2020): Fossil evidence for a Cretaceous rise of the mahogany family. Free access, American Journal of Botany, 107: 139-147. See also here (Science Daily), and there (PDF file, PhysOrg).

L. Augusto et al. (2014): The enigma of the rise of angiosperms: can we untie the knot? In PDF, Ecology Letters.

Francisco J. Ayala et al. (2000): Variation and evolution in plants and microorganisms: Toward a new synthesis 50 years after Stebbins. PNAS, 97: 6941-6944. Scroll to: "Trends and Patterns in Plant Evolution".

R.M. Bateman (2020): Hunting the Snark: the flawed search for mythical Jurassic angiosperms. In PDF, Journal of Experimental Botany, 71: 22–35. See also here.

J.M. Beaulieu et al. (2015): Heterogeneous rates of molecular evolution and diversification could explain the Triassic age estimate for angiosperms. Abstract.

J.M. Beaulieu et al. (2013): A Southern Hemisphere origin for campanulid angiosperms, with traces of the break-up of Gondwana. In PDF, BMC Evolutionary Biology, 13.

C.M. Belcher and V.A. Hudspith (2017): Changes to Cretaceous surface fire behaviour influenced the spread of the early angiosperms. New Phytologist, 213: 1521–1532.

! C.D. Bell et al. (2010): The age and diversification of the angiosperms re-revisited. In PDF, American Journal of Botany, 97: 1296-1303.

Museum of Paleontology, University of California (UCMP), Berkeley, CA: Introduction to the Anthophyta, Monocots versus Dicots, and Introduction to the Liliopsida. The Monocots.

J. Blanchard et al. (2016): Fruits, seeds and flowers from the Bovay and Bolden clay pits (early Eocene Tallahatta Formation, Claiborne Group), northern Mississippi, USA. In PDF, Palaeontologia Electronica. See also here.

! W.J. Bond and A.C. Scott (2010): Fire and the spread of flowering plants in the Cretaceous. In PDF, New Phytologist, 188: 1137-1150.

C. Kevin Boyce et al. (2010): Angiosperms Helped Put the Rain in the Rainforests: The Impact of Plant Physiological Evolution on Tropical Biodiversity. PDF file, Annals of the Missouri Botanical Garden, 97: 527-540.
Provided by the Internet Archive´s Wayback Machine.

C. Kevin Boyce et al. (2009). Angiosperm leaf vein evolution was physiologically and environmentally transformative. PDF file, Proceedings of the Royal Society B, 276: 1771-1776.

The palaeofiles. Articles here have all been prepared by students on the palaeobiology programmes in Bristol:
! Purported Triassic angiosperms.
Now provided by the Internet Archive´s Wayback Machine.

The palaeofiles. Articles here have all been prepared by students on the palaeobiology programmes in Bristol:
! The origin and evolution of angiosperms.
Now provided by the Internet Archive´s Wayback Machine.

T.J. Brodribb and T.S. Feild (2010): Leaf hydraulic evolution led a surge in leaf photosynthetic capacity during early angiosperm diversification. In PDF, Ecology Letters, 13: 175-183. See also here.
"... Our data suggest that early terrestrial angiosperms produced leaves with low photosynthetic rates, but that subsequent angiosperm success is linked to a surge in photosynthetic capacity during their early diversification".

G.E. Budd et al. (2021): The use of geological and paleontological evidence in evaluating plant phylogeographic hypotheses in the Northern Hemisphere Tertiary. Free access, See also here (in PDF).

Benjamin Burger, Utah State University, Vernal, Utah:
Why study fossil plants?
Invertebrate Paleontology and Paleobotany.
How did plants colonize the land, based on the fossil record?
How did the first seed plants (the Gymnosperms) evolve?
How did gymnosperms diversify during the early Mesozoic to become a modern dominate plant group?
How good is the fossil record of Cycads?
What is the significance of the fossil record of Ginkgo?
What is the fossil record of Horsetails?
Fossil Algae.
! What is an Angiosperm?
Video lectures.

! R.J. Burnham (2009): An overview of the fossil record of climbers: bejucos, sogas, trepadoras, lianas, cipós, and vines. PDF file, Rev. bras. paleontol., 12: 149-160.
Snapshot provided by the Internet Archive´s Wayback Machine.

R.J. Burnham (1994): Patterns in tropical leaf litter and implications for angiosperm paleobotany. In PDF, Review of Palaeobotany and Palynology.

R.J. Butler et al. (2009): Diversity patterns amongst herbivorous dinosaurs and plants during the Cretaceous: implications for hypotheses of dinosaur/angiosperm co-evolution. PDF file, Journal of Evolutionary Biol., 22: 446-459. See also here (abstract).

! A.S. Chanderbali et al. (2016): Evolving Ideas on the Origin and Evolution of Flowers: New Perspectives in the Genomic Era. In PDF, Genetics, 202: 1255–1265. See also here.

G. Chomicki et al. (2017): Evolution and ecology of plant architecture: integrating insights from the fossil record, extant morphology, developmental genetics and phylogenies. Annals of Botany 120: 855–891. See also here (in PDF).
Note Fig. 9: A timeline of plant architectures and branching mechanisms through time.

! Michael Clayton, Department of Botany, University of Wisconsin, Madison: Instructional Technology (BotIT). Some image collections. Excellent! Go to:

C. Coiffard et al. (2012): Rise to dominance of angiosperm pioneers in European Cretaceous environments , Abstract. See also here ( and there (

C. Coiffard et al. (2012): Deciphering Early Angiosperm Landscape Ecology Using a Clustering Method on Cretaceous Plant Assemblages. In PDF.

! M. Coiro et al. (2019): How deep is the conflict between molecular and fossil evidence on the age of angiosperms? Free access, New Phytologist, doi: 10.1111/nph.15708.
"... Critical scrutiny shows that supposed pre-Cretaceous angiosperms either represent other plant groups or lack features that might confidently assign them to the angiosperms. ..."

B. Cornet (1989): The reproductive morphology and biology of Sanmiguelia lewisii, and its bearing on angiosperm evolution in the Late Triassic. Evolutionary trends in Plants.

Bruce Cornet, (?) Raritan Valley Community College, Somerville, NJ: Why do Paleobotanists Believe in a Cretaceous Origin of Angiosperms? A controversial topic. This website presents palaeobotanical evidence on the origin of flowering plants, with evidence for and against a Cretaceous origin.

! P.R. Crane and A.B. Leslie (2013): Major Events in the Evolution of Land Plants. In PDF. The Princeton Guide to Evolution.
1. Phylogenetic framework.
2. Origin and diversification of land plants.
3. Origin and diversification of vascular plants.
4. Origin and diversification of seed plants.
5. Origin and diversification of flowering plants.
6. Innovation in the land plant body.
7. Innovation in land plant reproduction.
8. Co-evolution with animals.
9. Patterns of extinction.
See also here, and there (Google books).

P.R. Crane et al. (2010): Darwin and the Evolution of Flowers. PDF file, Phil. Trans. R. Soc. B, 365: 347-350.

! W.L. Crepet and K.J. Niklas (2009): Darwin´s second "abominable mystery": Why are there so many angiosperm species? PDF file, American Journal of Botany, 96: 366-381. See also here (abstract).

! W.L. Crepet (2008): The Fossil Record of Angiosperms: Requiem or Renaissance? Abstract, Annals of the Missouri Botanical Garden.


! W.L. Crepet et al. (2004): Fossil evidence and phylogeny: the age of major angiosperm clades based on mesofossil and macrofossil evidence from Cretaceous deposits. In PDF, American Journal of Botany, 91: 1666-1682.

William L. Crepet, Department of Plant Biology, Cornell University, Ithaca, NY: Progress in understanding angiosperm history, success, and relationships: Darwin's abominably "perplexing phenomenon". PNAS 2000; 97: 12939-12941.

Judith L. Croxdale, Department of Botany, University of Wisconsin, Madison (website hosted by Biology Online): Stomatal patterning in angiosperms. Stomatal pattern types, means of measuring them, advantages of each type of measurement, and then present patterning from evolutionary, physiological, ecological, and organ views are discussed.
Website outdated, a version archived by the Internet Archive Wayback Machine.

Charles Daghlian (Dartmouth College, Hannover, NH) and Jennifer Svitko, Paleobotanical Holdings at the Liberty Hyde Bailey Hortorium at Cornell University: Paleoclusia 3D Reconstructions. Movies from CT scans done on the Turonian fossils. See also here (W.L. Crepet and K.C. Nixon 1998, abstract and photos).
Snapshots provided by the Internet Archive´s Wayback Machine.

H.J. de Boer et al. (2012): A critical transition in leaf evolution facilitated the Cretaceous angiosperm revolution. In PDF, Nature Communications, 3. See also here.

! D.L. Dilcher (2001): Paleobotany: some aspects of non-flowering and flowering plant evolution. In PDF, Taxon.
For early angiospermous fossil floras see fig 1 (on PDF page 4).

David Dilcher (2000): Toward a new synthesis: Major evolutionary trends in the angiosperm fossil record. PDF file, Proc Natl Acad Sci U S A., 97: 7030-7036. See also here.

P. Donoghue (2019): Evolution: The Flowering of Land Plant Evolution. Abstract, Current Biology. See also:
Evolution: The flowering of land plant evolution - whence and whither? (in PDF).

Michael J. Donoghue and James A. Doyle (2000): Seed plant phylogeny: Demise of the anthophyte hypothesis? Current Biology, 10: R106-R109.

Stephen R. Downie and Kenneth R. Robertson, Life Sciences at the University of Illinois, Urbana-Champagne: Systematics of Plants. This course introduces the principles and methods of identifying, naming, and classifying flowering plants. It includes a survey of selected flowering plant families and provides information on their interrelationships. Go to: Digital Flowers.
These expired links are available through the Internet Archive´s Wayback Machine.

J.A. Doyle and P.K. Endress (2014): Integrating Early Cretaceous Fossils into the Phylogeny of Living Angiosperms: ANITA Lines and Relatives of Chloranthaceae Int. J. Plant Sci., 175: 555–600. See also here.

! J.A. Doyle (2012): Molecular and fossil evidence on the origin of angiosperms. In PDF, Annual Review of Earth and Planetary Sciences, 40: 301-26.

! J.A. Doyle (1998): Molecules, morphology, fossils, and the relationship of angiosperms and Gnetales. In PDF, Molecular phylogenetics and evolution.

M. Eberlein (2015): Bestimmungs- und Verbreitungsatlas der Tertiärflora Sachsens – Angiospermenblätter und Ginkgo. PDF file (in German). Thesis, University of Dresden. First part of a reference book of the Tertiary flora of Saxony. See also here (abstract).

! Peter K. Endress (2011): Angiosperm ovules: diversity, development, evolution. In PDF, Annals of Botany, 107: 1465-1489.

P.K. Endress and J.A. Doyle (2009): Reconstructing the ancestral angiosperm flower and its initial specializations. PDF file, American Journal of Botany, 96: 22-66.

! Mike Farabee, Estrella Mountain Community College Center, Avondale, Arizona:
On-Line Biology Book. Introductory biology lecture notes. Go to:
Flower Structure.
Fertilization and Fruits.
These expired links are available through the Internet Archive´s Wayback Machine.

Jeffrey A. Fawcett and Yves Van de Peer (2010): Angiosperm polyploids and their road to evolutionary success. Trends in Evolutionary Biology.

T.S. Feild et al. (2011): Fossil evidence for Cretaceous escalation in angiosperm leaf vein evolution. In PDF, PNAS, 108: 8363-8366.

Taylor S. Feild and N.C. Arens (2007): The ecophysiology of early angiosperms. PDF file, Plant, Cell and Environment, 30: 291-309.

T.S. Feild et al. (2004): Dark and disturbed: a new image of early angiosperm ecology. PDF file, Paleobiology, 30: 82-107.

S. Fields (2021): Diversification of Angiosperms During the Cretaceous Period. In PDF, Undergraduate Student Theses, Environmental Studies Program at DigitalCommons@University of Nebraska,Lincoln. See also here.

! C.S.P. Foster (2016): The evolutionary history of flowering plants. In PDF, Journal & Proceedings of the Royal Society of New South Wales, 149: 65-82.

William Friedman et al., Department of Ecology and Evolutionary Biology, University of Colorado, Boulder: Molecular and Organismal Research in Plant History, MORPH. MORPH, an NSF research coordination network, fosters cross-disciplinary interactions between organismic and molecular plant biologists studying the evolution of morphological diversity to promote a modern synthesis in plant evolutionary developmental biology. See also: Publications.
A version archived by the Internet Archive´s Wayback Machine.

! E.M. Friis et al. (2019): The Early Cretaceous Mesofossil Flora of Torres Vedras (Ne of Forte Da Forca), Portugal: A Palaeofloristic Analysis of an Early Angiosperm Community. Open access, Fossil Imprint, 75: 153–257. See also here (in PDF).
"... the oldest mesofossil flora containing angiosperm remains to be described in detail based on well-preserved flower, fruit and seed remains. It provides the most detailed information currently available on the structural diversity of angiosperms at this early stage in their evolution, the range of angiosperm species present, and their relationships to extant angiosperm lineages. ..."

E.M. Friis et al. (2015): Exceptional preservation of tiny embryos documents seed dormancy in early angiosperms. In PDF, Nature, 528: 551-554. See also here.

! E.M. Friis et al. (2011): Early Flowers and Angiosperm Evolution. Abstract, Cambridge University Press.
See also here (in PDF, long download time) and there (Google books).
Also worth checking out: Book Review, by P.J. Rudall, Botanical Journal of the Linnean Society, 170. In PDF.
"... This long-awaited book represents not only a remarkable tour de force of palaeobotanical literature, but also a potentially enduring biological textbook. ..."

Else Marie Friis, Swedish Museum of Natural History, Stockholm: Cretaceous angiosperms from Europe and North America (Silvianthemum suecicum), and Cretaceous angiosperms from Kazakhstan.
Snapshots taken by the Internet Archive´s Wayback Machine.

! Else Marie Friis et al. (2010): Diversity in obscurity: fossil flowers and the early history of angiosperms. PDF file, Phil. Trans. R. Soc. B, 365: 369-382. Some of the specimens are charcoalified and have retained their original three-dimensional shape. See also here.

! E.M. Friis, K. Raunsgaard Pedersen and Peter R. Crane (2006): Cretaceous angiosperm flowers: Innovation and evolution in plant reproduction. PDF file, Palaeogeography, Palaeoclimatology, Palaeoecology. 232 (2-4): 251-293.

! M.W. Frohlich & M.W. Chase (2007): After a dozen years of progress the origin of angiosperms is still a great mystery. In PDF, Nature 450: 1184-1189. See also here and there.

Q. Fu et al. (2018): An unexpected noncarpellate epigynous flower from the Jurassic of China. In PDF, eLife, 7: e38827. See also:
D.W. Taylor and H. Li (2019): Paleobotany: Did flowering plants exist in the Jurassic period.

Q. Fu et al. (2017): Nanjinganthus: An Unexpected Flower from the Jurassic of China. In PDF, bioRxiv (pronounced "bio-archive"). See also here.

J.F. Genise et al. (2020): 100 Ma sweat bee nests: Early and rapid co-diversification of crown bees and flowering plants. Open access, PLoS ONE 15: e0227789.

B. Gomez et al. (2015): Montsechia, an ancient aquatic angiosperm. In PDF, PNAS, 112: 10985–10988. See alao here.
Note Fig. 3: Reconstructions of Montsechia vidalii.

! R. Gorelick (2001): Did insect pollination cause increased seed plant diversity? PDF file, Biological Journal of the Linnean Society, 74: 407-427.

! G. Han et al. (2016): A Whole Plant Herbaceous Angiosperm from the Middle Jurassic of China. In PDF, Acta Geologica Sinica. See also here (abstract) and there (in German, with photograph and reconstruction).

Ben Harder, National Geographic News: Dino-Era Fossil—The First Flower? About the Archaefructaceae.

! P.S. Herendeen et al. (2017): Palaeobotanical redux: revisiting the age of the angiosperms. In PDF, Nature Plants 3. See also here.

P.A. Hochuli and S. Feist-Burkhardt (2013): Angiosperm-like pollen and Afropollis from the Middle Triassic (Anisian) of the Germanic Basin (Northern Switzerland). In PDF, Frontiers in plant science.

P.A. Hochuli, Paläontologisches Institut und Museum, Universität Zürich: Timing of angiosperm evolution. Research project description.

Natalia Holden, Department of Biological Sciences, University of Alberta, Edmonton, Canada: The early Angiosperms: Paleophytogeography and Depositional Settings. A slideshow.
This expired link is available through the Internet Archive´s Wayback Machine.

! S. Hu et al. (2008): Early steps of angiosperm-pollinator coevolution. PDF file, PNAS, 105: 40-245. See also here (abstract).

! Norman F. Hughes (1994): The Enigma of Angiosperm Origins. 405 pages. Provided by Cambridge University Press through the Google Print Publisher Program.

Norman F. Hughes (1982): Palaeobiology of Angiosperm Origins: Problems of Mesozoic Seed-Plant Evolution. Provided by Google books.

Marty Huss: Angiosperm Life Cycle. Powerpoint presentation.

N.A. Jud et al. (2018): A new fossil assemblage shows that large angiosperm trees grew in North America by the Turonian (Late Cretaceous). In PDF, Sci. Adv., 4: eaar8568.
"A large silicified log (maximum preserved diameter, 1.8 m; estimated height, ca. 50 m) is assigned to the genus Paraphyllanthoxylon; it is the largest known pre-Campanian angiosperm and the earliest documented occurrence of an angiosperm tree more than 1.0 m in diameter."

N.A. Jud (2015): Fossil evidence for a herbaceous diversification of early eudicot angiosperms during the Early Cretaceous. In PDF, Proc. R. Soc., B, 282. See also here.

N.A. Jud and L.J. Hickey (2013): Potomacapnos apeleutheron gen. et sp. nov., a new Early Cretaceous angiosperm from the Potomac Group and its implications for the evolution of eudicot leaf architecture. In PDF, Am. J. Bot., see also here.

! O. Katz (2018): Extending the scope of Darwin’s ‘abominable mystery’: integrative approaches to understanding angiosperm origins and species richness. Open access, Annals of Botany, 121: 1–8.
See also here (Botany One).

A.J. Kerkhoff et al. (2014): The latitudinal species richness gradient in New World woody angiosperms is consistent with the tropical conservatism hypothesis. In PDF, PNAS, 111: 8125–8130. See also here.

! Kimball´s Biology Pages (by John W. Kimball). The pages represent an online biology textbook. Go to: Sexual Reproduction in Angiosperms.

Michael Knee, Department of Horticulture and Crop Science, Ohio State University, Columbus: General Plant Biology Online Resources. A version archived by Internet Archive Wayback Machine. Lecture notes. Go to: Evolution and diversity of angiosperms.

Knox College, Galesburg, Illinois: The Seed Plants: Gymnosperms & Angiosperms. Powerpoint presentation.

Ari and Susan Kornfeld, Natural Perspective: The Plant Kingdom: Dicots Overview. A version archived by Internet Archive Wayback Machine.

V. Krassilov and S. Barinova (2014): "Flower" of Magnolia grandiflora is not flower and what about "basal angiosperms". In PDF, Journal of Plant Sciences, 2: 282-293.

V. Krassilov (2012): Fossil record of angiosperm origin: new evidence and interpretation. In PDF, Horizons in Earth Science Research, 8. (Nova Publishers, New York).

V.A. Krassilov (2009): Diversity of Mesozoic Gnetophytes and the First Angiosperms. PDF file, Paleontological Journal, 43: 1272-1280. A version archived by Internet Archive Wayback Machine.

A. Kremer and A.L. Hipp (2019): Oaks: an evolutionary success story. Free access, New Phytologist, doi: 10.1111/nph.16274.

B.B. Lamont and T. He (2012): Fire-adapted Gondwanan Angiosperm floras evolved in the Cretaceous. In PDF, BMC Evolutionary Biology, 12. See also here.

Gerhard Leubner Lab, University Freiburg, Germany: Seed Evolution. Go to: Angiosperm seed evolution and species diversification.

Gerhard Leubner, The Seed Biology Place, Molecular Plant Sciences, University Freiburg, Germany: Seed evolution. Origin and evolution of the seed habit. See also: Seed dictionary English-German.

Z.-J. Liu and X. Wang (2017): Yuhania: a unique angiosperm from the Middle Jurassic of Inner Mongolia, China. Open access, Historical Biology, 29: 431-441.

! H.T. Li et al. (2019): Origin of angiosperms and the puzzle of the Jurassic gap. Abstract, Nature Plants, 5: 461–470. See also here (in PDF).
"... With a well-resolved plastid tree and 62?fossil calibrations, we dated the origin of the crown angiosperms to the Upper Triassic, with major angiosperm radiations occurring in the Jurassic and Lower Cretaceous. This estimated crown age is substantially earlier than that of unequivocal angiosperm fossils, and the difference is here termed the ‘Jurassic angiosperm gap’. ..."

Biological Sciences, Ohio State University, Lima: Plant Biology at OSU Lima.
This expired link is now available through the Internet Archive´s Wayback Machine.

A. Linkies et al. (2010): The evolution of seeds. PDF file, New Phytologist.

S.A. Little et al. (2014): Reinvestigation of Leaf Rank, an Underappreciated Component of Leo Hickey´s Legacy. In PDF.

Z.J. Liu et al. (2019): Zhangwuia: an enigmatic organ with a bennettitalean appearance and enclosed ovules. In PDF, Earth and Environmental Science Transactions of The Royal Society of Edinburgh, 108 (Agora Paleobotanica), : 419-428. See also here.

Z.-J. Liu et al. (2018): The Core Eudicot Boom Registered in Myanmar Amber. Open access, Scientific Reportsvolume 8.
Note figure 5: Reconstruction of Lijinganthus revoluta.

Z.-J. Liu and X. Wang (2016): A perfect flower from the Jurassic of China. In PDF, Historical Biology, 28: 707-719. See also here (Abstract).

W.-Z. Liu et al. (2014): From leaf and branch into a flower: Magnolia tells the story. In PDF, Botanical Studies, 55.

Department of Botany, University of Wisconsin, Madison: Plant Systematics Collection. This web site provides structured access to a teaching collection of plant images representing over 250 families and 1000 genera of vascular plants. Go to: Phylum Magnoliophyta (Flowering Plants).

S. Magallón et al. (2015): A metacalibrated time-tree documents the early rise of flowering plant phylogenetic diversity. In PDF, New Phytologist.

! S. Magallón (2009): Flowering plants (Magnoliophyta). PDF file, In: S.B. Hedges and S. Kumar (eds.): The Timetree of Life (see here).

Susana Magallón and Amanda Castillo (2009): Angiosperm diversification through time. PDF file, American Journal of Botany, 96: 349-365.

Steven R. Manchester (website hosted by International Organisation of Palaeobotany): Living Fossils, Davidia - the Dove Tree and its fossil record.

! J. Massoni et al. (2014): Fossil calibration of Magnoliidae, an ancient lineage of angiosperms. Palaeontologia Electronica.

Duane D. McKenna et al. (2009): Temporal lags and overlap in the diversification of weevils and flowering plants. PDF file, PNAS, 106: 7083-7088. See also here (abstract).

Department of Biological Sciences, University of Memphis: History of Angiosperm Classification (Post Darwin´s Theory of Evolution). In PDF.

! John M. Miller (, University of California, Berkeley: Origin of Angiosperms. See also here or navigate from essay contents.

! Sebastian Molnar, Department of Zoology, University of British Columbia, Vancouver: Evolution and the Origins of Life. A directory of introductions concerning evolution, with a bias to Plant Biology and Evolution. Go to: Angiosperm Origins and Evolution.

Nature Science Update (December 8, 1999): One for the Vine. "A prickly climbing vine", Vasovinea tianii (Gigantopteridales), that lived more than 250 million years ago could shed light on the origin of flowering plants.
This expired link is available through the Internet Archive´s Wayback Machine.

D.L. Nickrent (2020): Parasitic angiosperms: How often and how many? Free access, Taxon.

! Dan Nickrent, Kevin C. Nixon & Dale Vitt (Curatorial Board; website served from Southern Illinois University, Carbondale): Phyto Images. This site includes a wide variety of vascular plant and bryophyte photos of high quality. Phyto Images belongs to DOL (, which is a web interface based on the Encino Software Project. The Encino project is a unified set of software tools for storing, retrieving, and analyzing biodiversity. Search the Cladogramm Database or use the Diagnostic Keys. Superbly done!

A.B. Nicotra et al. (2011): The evolution and functional significance of leaf shape in the angiosperms. In PDF, Functional Plant Biology, 38: 535-552.

NOVA (science series on television): First Flower.

Daniel Oakley et al. (2009): Morphometric analysis of some Cretaceous angiosperm woods and their extant structural and phylogenetic analogues: Implications for systematics. PDF file, Review of Palaeobotany and Palynology, 157: 375-390.

J. Ollerton and E. Coulthard (2009): Evolution of Animal Pollination. In PDF, Science, 326.

! D. Peris et al. (2017): False Blister Beetles and the Expansion of Gymnosperm-Insect Pollination Modes before Angiosperm Dominance. In PDF, Current Biology, 27. See also here.

Marc Philippe et al. (2008): Woody or not woody? Evidence for early angiosperm habit from the Early Cretaceous fossil wood record of Europe. PDF file, Palaeoworld, 17: 142-152.

Ray Phillips, Information Technology Services, Colby College, Waterville, Maine: World Wide Flowering Plant Family Identification. Select the characters that are present in the specimen being identified and press "Submit". Database is part of "Biology 211: Flowering Plant Taxonomy", an introduction to the principles and practice of flowering plant taxonomy.

Tõnu Ploompuu, Biology, Tallinn Pedagogical University, Tallinn, Estonia: Resting and active evolution. Possible preadaptations in the early evolution of Angiosperms. See also here.

George Poinar and Greg Poinar (2018): The antiquity of floral secretory tissues that provide today’s fragrances. Abstract, Historical Biology. See also:
Schnupperten schon Dinos Blumenduft? Kreidezeitliche Blütenpflanzen könnten bereits Düfte produziert haben In German,

G. Poinar et al. (2016): Fossil species of Boehmerieae Gaudich. (Urticaceae) in Dominican and Mexican amber: A new genus (Ekrixanthera) and two new species with anemophilous pollination by explosive pollen release, and possible lepidopteran herbivory. In PDF, Botany.

D. Pons and D. de Franceschi (2007): Neogene woods from western Peruvian Amazon and palaeoenvironmental interpretation. Bulletin of Geosciences, 82: 343-354.

I. Poole (2000): Fossil angiosperm wood anatomy: its role in the reconstruction of biodiversity and palaeoenvironment. PDF file, Botanical journal of the Linnean Society, 134: 361-381.

H. Prier et al. (2004): Exotische Gehölze im KIRCHHEIMER-Arboretum Freiburg. PDF file, in German. LGRB-Informationen, Heft 15 (Landesamt für Geologie, Rohstoffe und Bergbau Baden-Württemberg, Freiburg i. Br.). See also here.

K.M. Pryer et al. (2001): Horsetails and ferns are a monophyletic group and the closest living relatives to seed plants. Abstract, Nature, 409: 618-622.
! See also here (in PDF).

Public Broadcasting Service (PBS): NOVA (a high rated science series on TV). First Flower. Go to: Flowers Modern and Ancient. About Archaefructus liaoningensis, discovered in the fossil beds of Liaoning Province in northeastern China.

! J.-H. Ran et al. (2018): Phylogenomics resolves the deep phylogeny of seed plants and indicates partial convergent or homoplastic evolution between Gnetales and angiosperms. Abstract.

! T. Reichgelt et al. (2018): The relation between global palm distribution and climate. Free access, Scientific Reports, 8:4721, doi:10.1038/s.

D. Ren et al. (20099. A Probable Pollination Mode Before Angiosperms: Eurasian, Long-Proboscid Scorpionflies. In PDF, Science, 326: 840-847. See also here.

James L. Reveal, Norton-Brown Herbarium, University of Maryland: Advanced Plant Taxonomy. Systems of classification for magnoliophyta, history of systematic botany, approaches to biological classification, taxonomic hierarchy, types of data.

Anita Roth-Nebelsick et al. (2001): Evolution and Function of Leaf Venation Architecture: A Review. PDF file, Annals of Botany 87: 553-566. See also here.

Gar W. Rothwell, Department of Environmental and Plant Biology, Ohio University, Athens: Angiophytes: Using Whole Plant Concepts to Interpret Angiosperm Origins.
The link is to a version archived by the Internet Archive´s Wayback Machine.
Selected Literature.

Gar W. Rothwell, Department of Environmental and Plant Biology, Ohio University, Athens, OH: Vascular Plant Morphology. A version archived by Internet Archive Wayback Machine. This course covers the structure, development, reproductive biology and relationships of vascular plants. The course is structured to emphasize the evolutionary changes that led to the diversity of modern tracheophytes. Go to Flowering Plants (PDF file).

D. Royer et al. (2010): Leaf economic traits from fossils support a weedy habit for early angiosperms. PDF file, American Journal of Botany, 97: 438-445. See also here, and there.

! Dmitry A. Ruban (2012): Mesozoic mass extinctions and angiosperm radiation: does the molecular clock tell something new? In PDF, Geologos, 18: 37-42.

P.J. Rudall and R.M. Bateman (2019): Leaf surface development and the plant fossil record: stomatal patterning in Bennettitales. Abstract, Biological Reviews.
"... Fossil bennettites – even purely vegetative material – can be readily identified by a combination of epidermal features, including distinctive cuticular guard-cell thickenings, lobed abaxial epidermal cells (‘puzzle cells’), transverse orientation of stomata perpendicular to the leaf axis, and a pair of lateral subsidiary cells adjacent to each guard-cell pair (termed paracytic stomata). ..."

P.J. Rudall et al. (2017): Evolution and development of monocot stomata. In PDF, American journal of botany, 104: 1122-1141.

Paula J. Rudall and Richard M. Bateman (2010): Defining the limits of flowers: the challenge of distinguishing between the evolutionary products of simple versus compound strobili. In PDF, Philos. Trans. R. Soc. London, B Biol. Sci., 365: 397-409. See also here (abstract).

A. Salt (2019): When did the first flowers open? Botany One.

! H. Sauquet and S. Magallón (2018): Key questions and challenges in angiosperm macroevolution. In PDF, New Phytologist, 219: 1170–1187. See also here.

! H. Sauquet et al. (2017): The ancestral flower of angiosperms and its early diversification. Nature Communications, 8.

S. Sgorbati et al. (2018): Was Charles Darwin right in his explanation of the ‘abominable mystery’? Free access, Italian Botanist, 5: 25–30.

G. Shi et al. (2021): Mesozoic cupules and the origin of the angiosperm second integument, Abstract, Nature, 594: 23–226. See also here (in PDF).

! D. Silvestro et al. (2020): Fossil data support a pre-Cretaceous origin of flowering plants. In PDF, Nature Ecology & Evolution. See also here.
"... Yet, our results indicate that an early, pre-Cretaceous origin of angiosperms is supported not only by molecular phylogenetic hypotheses but also by an analysis of the fossil record ..."

! Stephen A. Smith et al. (2010): An uncorrelated relaxed-clock analysis suggests an earlier origin for flowering plants. PDF file, PNAS, 107: 5897-5902. See also here, or there.

! S.A. Smith and M.J. Donoghue (2008): History in Flowering Plants Rates of Molecular Evolution Are Linked to Life. In PDF, Science, 322. See also here (abstract).

D.D. Sokoloff et al. (2019): Supposed Jurassic angiosperms lack pentamery, an important angiosperm-specific feature. Open access, New Phytologist.

D. Soltis et al. (2017): Phylogeny and Evolution of the Angiosperms. Book announcement. See also here (Google books). Worth checking out:
! Relationships of Angiosperms to Other Seed Plants. In PDF.

P.S. Soltis et al. (2019): Darwin review: angiosperm phylogeny and evolutionary radiations. In PDF, Proc. R. Soc. B, 286: 20190099. See also here.

! D.E. Soltis et al. (2009): Polyploidy and angiosperm diversification. In PDF, Am. J. Bot., 96: 336-348.

Pamela Soltis, (a resource of the American Institute of Biological Sciences): Flowering Plants: Keys to Earth´s Evolution and Human Well-Being. A version archived by Internet Archive Wayback Machine.

Pamela S. Soltis and Douglas E. Soltis (2004): The origin and diversification of angiosperms. PDF file, American Journal of Botany, 91: 1614-1626.

Space Daily: Paleobotanist identifies what could be the mythical "first flower". (August 18, 2015).

! Doug Soltis, Amber Tilley and Hongshan Wang, Florida Museum of Natural History (FLMNH), University of Florida: Deep Time. A comprehensive phylogenetic tree of living and fossil angiosperms. Deep Time explore the ways in which angiosperm fossils can be appropriately integrated into the phylogenetic framework for extant taxa, with the ultimate goal of forming a comprehensive phylogenetic tree of living and fossil angiosperms. This includes the evaluation and prioritization of the fossil record, the critical appraisal of the age of fossils, the construction of a morphological data matrix for fossils and extant angiosperms, the integration of fossils into the angiosperm tree and the calibration of divergence times.

! P.F. Stevens and Hilary Davis, Missouri Botanical Garden, St. Louis: Angiosperm Phylogeny Website. The focus of this site is on angiosperms and emphasis is placed on plant families. You can also navigate from the Orders- or the Families-website. Go to:

! P.F. Stevens and Hilary Davis, Missouri Botanical Garden, St. Louis: Angiosperm Phylogeny Website. The focus of this website is about classification on angiosperm plant families. Sets of evolutionary trees grouping families and orders and showing details of the arrangement. Don´t miss the complete synonymy of family name directory and some literature references. Excellent!

Ruth A. Stockey, Department of Biological Sciences, University of Alberta, Edmonton: Paleobotany of Angiosperm Origins. Go to: Course Outline. Chiefly bibliographies and weblinks.
These expired links are now available through the Internet Archive´s Wayback Machine.

Ge Sun, David L. Dilcher, Shaoling Zheng, and Zhekun Zhou: In Search of the First Flower: A Jurassic Angiosperm, Archaefructus, from Northeast China. Science 1998: 1692-1695.
Snapshot provided by the Internet Archive´s Wayback Machine.

Syllabus of Plant Families, A. Engler´s Syllabus der Pflanzenfamilien (13th edition by Wolfgang Frey):
4 Pinopsida (Gymnosperms), Magnoliopsida (Angiosperms) p.p.: Subclass Magnoliidae [Amborellanae to Magnolianae, Lilianae p.p. (Acorales to Asparagales)].

Ralph E. Taggart, Department of Botany and Plant Pathology/Department of Geological Sciences at Michigan State University, East Lansing:
! BOT335 Lecture Schedule. Some interesting chapters in terms of palaeobotany, e.g.
The First Vascular Land Plants;
Carboniferous Forests;
Arborescent Lycopods;
Psaronius: a Carboniferous tree-fern;
Carboniferous Horsetails;
Carboniferous Seed Ferns;
The Evolution of Conifers;
Cycadophytes, the True Cycads;
Mesozoic Cycadeoids;
North American Redwoods, Past and Present.
These expired links are available through the Internet Archive´s Wayback Machine.

D.W. Taylor and H. Li (2018): Paleobotany: Did flowering plants exist in the Jurassic period? eLife, 7: e43421.
"... we infer that Nanjinganthus shows substantial similarity to predicted models of ancestral characters and Early Cretaceous angiosperms, so the evidence suggests that it is a Jurassic flowering plant. ..."

David Winship Taylor et al. (2006): Biogeochemical evidence for the presence of the angiosperm molecular fossil oleanane in Paleozoic and Mesozoic non-angiospermous fossils. Abstract, Paleobiology, 32: 179-190.

! David W. Taylor and Leo J. Hickey (1996): Flowering Plant Origin, Evolution & Phylogeny. Google books (some pages omitted); American Institute of Biological Sciences (Springer), 404 pages.

Edith L. Taylor and Thomas N. Taylor (2009): Seed ferns from the late Paleozoic and Mesozoic: Any angiosperm ancestors lurking there? PDF file, American Journal of Botany, 96: 237-251. See also here.

H. Hamshaw Thomas (1936): Palaeobotany and the origin of the angiosperms. PDF file, The Botanical Review, 2: 97-418.

Greg Thorn, Department of Biology, University of Western Ontario, Canada:
! Evolution of Plants . Lecture notes, e.g.:
Evolution of the Angiosperms. Powerpoint presentation.

Greg Thorn, Department of Biology, University of Western Ontario: Evolution of Plants (Powerpoint presentations). Navigate from here with information from the Syllabus. See e.g. Lecture 16: Evolution of Plants. The evolution of early angiosperms.

! B.H. Tiffney and S.R. Manchester (2001): The Use of Geological and Paleontological Evidence in Evaluating Plant Phylogeographic Hypotheses in the Northern Hemisphere Tertiary. Abstract, International Journal of Plant Sciences, 162. See also here (in PDF).

! Amber Tilley and Hongshan Wang, Florida Museum of Natural History, University of Florida: Deep Time. Projects about the origin of angiosperms.

Tree of Life Web Project: Angiosperms (by Pam Soltis, Doug Soltis, and Christine Edwards).

G.R. Upchurch Jr. (1984): Cuticle evolution in Early Cretaceous angiosperms from the Potomac Group of Virginia and Maryland. PDF file, Annals of the Missouri Botanical Garden. A version archived by Internet Archive Wayback Machine.

! C.J. van der Kooi and J. Ollerton (2020): The origins of flowering plants and pollinators. Free access, Science, 368: 1306-1308.
See also here (in PDF).

S. Wang et al. (2021): Cretaceous fire-resistant angiosperms. In PDF, preprint, DOI:
See also here.
"... both preserved as inclusions in Cretaceous amber from northern Myanmar (~99 Ma). These specialized flower structures, named Phylica piloburmensis sp. nov. and Eophylica priscastellata gen. et sp. nov., were adapted to surviving frequent wildfires, providing the earliest evidence of fire-resistance in angiosperms. ..."

H. Wang et al. (2013): Fruits, seeds, and flowers from the Warman clay pit (middle Eocene Claiborne Group), western Tennessee, USA. In PDF, Palaeontologia Electronica. See also here.

X. Wang: An Era of Errors. About Archaeanthus (in PDF). See also here.

! X. Wang (2017): A Biased, Misleading Review on Early Angiosperms. In PDF, Natural Science, 9: 399-405.
Please note: P.S. Herendeen et al. (2017): Palaeobotanical redux: revisiting the age of the angiosperms. In PDF, Nature Plants 3. See also here.

X. Wang (2010): Suggested Angiosperm Ancestors. In PDF, Chapter 2, The Dawn Angiosperms. Lecture Notes in Earth Sciences Volume 121: 5-16. See also here (Springer) and there Google books.

X. Wang et al. (2007): Schmeissneria: A missing link to angiosperms? Open access, BMC Evolutionary Biology, 7. See also here.

! L. Watson Albany, Australia, and M. J. Dallwitz CSIRO Entomology, Canberra, Australia (page hosted by DELTA): The Families of Flowering Plants. Descriptions, illustrations, identification, and information retrieval. Version: 25th November 2009. The Intkey software is required.
Also worth to check out: Static information. Character list, implicit attributes, notes on the APG classification, etc.
Snapshot taken by the Internet Archive´s Wayback Machine.

G.D.A. Werner et al. (2014): A single evolutionary innovation drives the deep evolution of symbiotic N2-fixation in angiosperms. Open access, Nature Communications, 5: 4087.

Biology Department, Western Washington University, Bellingham, Washington:
! Flowering vascular plants. Powerpoint presentation. See also here, or there.

E.A. Wheeler and S.R. Manchester (2007): Review of the wood anatomy of extant Ulmaceae as context for new reports of late Eocene Ulmus woods. PDF file, Bulletin of Geosciences, 82: 329-342.

Niklas Wikström et al. (2001): Evolution of the angiosperms: calibrating the family tree. PDF file, Proc. R. Soc. Lond., B, 268: 2211-2220.

! Wikipedia, the free-content encyclopedia: Spermatophyte. Go to: Flowering Plant, and Magnoliopsida.

P. Wilf (2008): Fossil angiosperm leaves: paleobotany´s difficult children prove themselves. PDF file, Paleontological Society Papers, 14: 319-333.

Kathy Willis and Jennifer McElwain: Flowering plant origins. Provided by the Internet Archive´s Wayback Machine.
In PDF, from: The Evolution of Plants (Oxford University Press).
Worth checking out: Second Edition.

Kathy Willis, School of Geography and the Environment, University of Oxford, & Jenny McElwain, Field Museum of Natural History, Chicago (Oxford University Press): The Evolution of Plants. Book announcement. Go to: Chapter 06, Flowering plant origins (PDF file).
Snapshots provided by the Internet Archive´s Wayback Machine.

! Hugh D. Wilson, Department of Biology Herbarium (TAMU), Texas A&M University: Flowering Plant Gateway. This project involves the development of computer programs that allow automated, machine-generated HTML page production for each Subclass/Superorder of the Flowering Plants as structured by the Cronquist, Takhtajan, and Thorne Systems of classification.

! S.L. Wing and L.D. Boucher (1998): Ecological aspects of the Cretaceous flowering plant radiation. In PDF, Annu. Rev. Earth Planet. Sci. 1998 26: 379-421.

! A.E. Zanne et al. (2014): Three keys to the radiation of angiosperms into freezing environments. In PDF, Nature. Provided by the Internet Archive´s Wayback Machine.

L. Zhang et al. (2020): The water lily genome and the early evolution of flowering plants. Open access, Nature, 577: 79–84. Worth checking out:
Fig. 1d: Summary phylogeny and timescale of 115 plant species. Blue bars at nodes represent 95% credibility intervals of the estimated dates.

X. Zhang et al. (2017): How the ovules get enclosed in magnoliaceous carpels. PLoS ONE, 12: e0174955.

Top of page
Links for Palaeobotanists
Search in all "Links for Palaeobotanists" Pages!
index sitemap advanced
site search by freefind

This index is compiled and maintained by Klaus-Peter Kelber, Würzburg,
Last updated October 29, 2021

eXTReMe Tracker